Mesozooplankton and Copepod Community Structures in the Southern East China Sea: the Status During the Monsoonal Transition Period in September
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Helgol Mar Res (2012) 66:621–634 DOI 10.1007/s10152-012-0296-1 ORIGINAL ARTICLE Mesozooplankton and copepod community structures in the southern East China Sea: the status during the monsoonal transition period in September Li-Chun Tseng • Hans-Uwe Dahms • Qing-Chao Chen • Jiang-Shiou Hwang Received: 14 April 2011 / Revised: 4 February 2012 / Accepted: 10 February 2012 / Published online: 25 February 2012 Ó Springer-Verlag and AWI 2012 Abstract A field sampling was conducted before the onset and comprising about 50% of all amphipods. Chaetognath of the northeasterly monsoon to investigate the copepod abundance showed a significantly negative correlation with community composition during the monsoon transition salinity (r = 0.77, p = 0.027), whereas mesozooplankton period at the northern coast of Taiwan (East China Sea). In group numbers had a significantly positive correlation with total, 22 major mesozooplankton taxa were found, with the salinity (r = 0.71, p = 0.048). Densities of four copepod Calanoida (relative abundance: 66.36%) and Chaetognatha species (Calanus sinicus, Calocalanus pavo, Calanopia (9.44%) being the most abundant. Mesozooplankton densi- elliptica and Labidocera acuta) showed a significantly neg- ties ranged between 226.91 and 2162.84 individuals m-3 ative correlation with seawater temperature. Communities of (mean ± SD: 744.01 ± 631.5 individuals m-3). A total of mesozooplankton and copepods of northern Taiwan varied 49 copepod species were identified, belonging to 4 orders, 19 spatially with the distance to land. The results of this study families, and 30 genera. The most abundant species were: provide evidence for the presence of C. sinicus in the coastal Temora turbinata (23.50%), Undinula vulgaris (17.92%), area of northern Taiwan during the early northeast monsoon and Acrocalanus gibber (14.73%). The chaetognath Flacc- transition period in September. isagitta enflata occurred at all 8 sampling stations, providing a 95% portion of the overall chaetognath contribution. Keywords Mesozooplankton Á Copepod Á Amphipoda were abundant at stations 4 and 5, with Hype- East China Sea Á Monsoon transition period rioides sibaginis and Lestigonus bengalensis being dominant, Introduction Communicated by Heinz-Dieter Franke. The marine biota in the waters around the island of Taiwan Li-Chun Tseng and Hans-Uwe Dahms are contributed equally to the is highly diverse. It has been estimated that around 10% of present contribution. the world’s marine species can be found in waters around L.-C. Tseng Á J.-S. Hwang (&) Taiwan (Shao 1998). Several authors explained this high Institute of Marine Biology, National Taiwan Ocean University, biodiversity as being caused by the convergence of several 2 Pei-Ning Road, Keelung 20224, Taiwan large water masses (Jan et al. 2002; Hwang et al. 2010a; e-mail: [email protected] Tseng et al. 2011a). Copepod communities in Taiwan L.-C. Tseng waters are highly influenced by water masses from ocean e-mail: [email protected] currents (Hwang et al. 2004, 2006, 2010b; Dur et al. 2007; H.-U. Dahms Tseng et al. 2008b, d, e; Lan et al. 2009). Temperate Green Life Science Department, College of Convergence, species are transferred from the north and tropical species Sangmyung University, Seoul 110-743, Korea from the south via the Taiwan Strait, affecting the meso- zooplankton communities on the west coast of Taiwan Q.-C. Chen South China Sea Institute of Oceanology, (Hwang et al. 2004, 2006, 2009; Dur et al. 2007; Tseng Chinese Academy of Science, Guangzhou, China et al. 2008a, e). 123 622 Helgol Mar Res (2012) 66:621–634 The Kuroshio Current flows from the equator via the Table 1 Locations and dates/day times of sampling on cruise ORII- east coast of the Philippines and brings warm water masses CR488 (2–3/September/1998) all along the east coast of Taiwan year round. In this Station Latitude Longitude Date manner, many warm water species from the south reach the (N) (E) (September/day time) western and northern coastal areas of Taiwan (Hsiao et al. 125°09.8090 121°46.5200 02/18:04 2004; Hwang et al. 2007; Tseng et al. 2008a, b, e). In the 225°14.9570 121°40.3550 02/19:17 Taiwan Strait and at the southern edge of the East China 325°21.7170 121°47.0540 02/20:59 Sea (ECS), water circulation is strongly influenced by the 425°27.6660 121°52.1050 02/22:32 prevailing monsoonal winds and their seasonal changes 525°33.5460 121°58.4690 03/00:25 (Lee and Chao 2003; Tseng et al. 2008b). During the 0 0 northeast (NE) monsoon period in winter and spring 625°39.874 122°04.830 03/02:00 0 0 (September to March), the China Coastal Current (CCC) 725°45.529 122°10.716 03/03:48 0 0 brings cold water masses from the Bohai Sea, the Yellow 825°51.256 122°16.918 03/06:00 Sea and the ECS to the Taiwan Strait (Wong et al. 2000; Liang et al. 2003; Lo et al. 2004a; Zuo et al. 2006; Tseng Taiwan. The sampling stations included one station near et al. 2008b). The southwest (SW) monsoon brings species Bi-Sar Harbor (northern Taiwan) and seven stations located from the South China Sea during summer and autumn on the transect between northern Taiwan and Pengjiayu Island (April to August) to the south of Taiwan (Lo et al. 2001; in the southern ECS, between 25°09.8090–25°51.2560Nand Liao et al. 2006; Tseng et al. 2008b; Lan et al. 2004, 2009). 121°40.3550–122°16.9180E(Table1, Fig. 1). Such complex water exchange affects the composition A research cruise was conducted in September 1998, i.e. and abundance of oceanic biota around the island of before the onset of the northeasterly monsoon. Zooplankton Taiwan. This also holds for microzooplankton (Vandromme samples were collected at the 8 selected stations by surface et al. 2010; Chang et al. 2011) and the mesozooplankton tows (0–5 m) with a standard North Pacific zooplankton net dominated by copepods (Liao et al. 1999; Tseng et al. (mouth diameter 45 cm, mesh size 333 lm) provided with a 2008e). Pelagic copepods are important food sources for fish flow meter in the center of the net opening (Hydrobios; Kiel, (Dahms and Hwang 2007; Mahjoub et al. 2011) and likely Germany). Samples were preserved immediately in 5% buf- play a pivotal role in the transfer of matter and energy in the fered formaldehyde in seawater. Prior to plankton collection, southern Taiwan Strait (Tseng et al. 2008c) and in the salinity and temperature were measured on board. In the northern South China Sea (Tseng et al. 2009). laboratory, samples were split by a Folsom splitter until the Zooplankton communities have long been applied as subsample contained less than 500 specimens. Copepods and indicators of water movements (Hwang and Wong 2005; mesozooplankton were sorted and identified at the species Dahms and Hwang 2010). In northern Taiwan, long-term level using the identification aids of Chen and Zhang (1965), studies of copepod successions showed that Calanus sinicus is Chen et al. (1974) and Chihara and Murano (1997). Additional an important indicator species for cold water masses from the original papers were used for species identification if required. Bohai Sea and the Yellow Sea (Hwang et al. 2004, 2006;Dur et al. 2007). Hwang et al. (2004)describedC. sinicus as rare in Statistical analyses samples from July to September, but the species appeared again in October, dramatically increasing in density and Species density matrices were analyzed using multivariate occurrence ratio. As yet, however, it is not known when analyses for copepod species. Non-metric cluster analysis exactly C. sinicus reaches the coastal areas of northern Taiwan was performed together with the Bray–Curtis similarity from the north. The present study aimed to characterize the index after logarithmic transformations of species abun- mesozooplankton community and copepod assemblages dance data. Significance levels of copepod assemblages during the southwest (SW)–northeast (NE) monsoon transi- between stations were calculated using a similarity pro- tion period in northern Taiwan waters because this appears to gram provided by the Plymouth Routine in Multivariate be an important period for community alterations. Ecological Research (PRIMER), version IV software package (Clarke and Warwick 1994). The abundance of copepod species of whole samples were used for the cal- Materials and methods culation of similarities before clustering. The functional test of Box and Cox (1964) for the transformation of data Field sampling was applied before the similarity analysis. The value (k)of the power transformation was set as 0.95, and the log For the present study, we selected the coastal waters of the (x ? 1) was applied to treat the individual densities of all southern ECS in the vicinity of Taipei County of northern samples. 123 Helgol Mar Res (2012) 66:621–634 623 Fig. 1 Map of the sampling stations on cruise ORII-CR488 during 2–3/September/1998 The copepod species characterizing each cluster were the IndVal indices. To evaluate copepod assemblages for further identified using the Indicator Value Index (IndVal) the present sampling period, an analysis of indicator spe- proposed by Dufreˆne and Legendre (1997). This index is cies was applied to each sampling cruise separately. obtained by multiplying the product of two independently The Shannon-Wiener diversity index was used to eval- computed values by 100: uate species diversity, and Pielou’s evenness was used to IndValðj; sÞ¼100SPðj; sÞFIðj; sÞ measure the relative abundance of species at each station. The relationship between copepod abundance and tem- where (SPj, s) is the specificity, and (FIj, s) is the fidelity of perature and salinity was studied with Pearson’s product a species (s) towards a group of samples (j), and these are moment correlation.